mammal

mammal

mammal, an animal of the highest class of vertebrates, the Mammalia. The female has mammary glands, which secrete milk for the nourishment of the young after birth. In the majority of mammals the body is partially or wholly covered with hair; the heart has four chambers, and only the left aortic arch is present; and a muscular diaphragm separates the chest from the abdominal cavity. Mammals are warm-blooded; that is, they have a relatively constant body temperature independent of the temperature of their surroundings. The mature red blood cells (erythrocytes) usually lack a nucleus. Except for the egg-laying monotremes (the platypus and the echidna, or spiny anteater), mammals give birth to live young. A marsupial is born in a more undeveloped state than the young of other mammals, although all are relatively helpless at birth. In some marsupials and in higher mammals the young receive prenatal nourishment through a placenta. The order Carnivora, or flesh-eating animals, includes terrestrial families such as the cat, dog, and bear as well as the aquatic seal, sea lion, and walrus. Other aquatic mammals are the whale, porpoise, and dolphin of the order Cetacea and the manatee and dugong of the order Sirenia. Unusual adaptations are also found in the bat (order Chiroptera); in the elephant (order Proboscidea); in the sloth, armadillo, and anteater (order Edentata); and in the beaver, woodchuck, porcupine, and squirrel (order Rodentia). The order Insectivora includes the shrew and the mole. There are two groups of ungulates, or hoofed mammals: most members of the order Perissodactyla, including the horse and the rhinoceros, are odd-toed, with the third digit the largest; those of the order Artiodactyla, including the deer, antelope, camel, pig, and cow, are even-toed, with the third and fourth digits symmetrical and functional. Humans, monkeys, apes, and lemurs belong to the order Primates. Some remains of mammals are identified as from the Jurassic period of the Mesozoic era; the group became diversified in the Tertiary period of the Cenozoic era.

All mammals (except for the five species of monotremes) give birth to live young instead of laying eggs. Most mammals also possess specialized teeth, and the largest group of mammals, the placentals, use a placenta during gestation. The mammalian brain regulates endothermic and circulatory systems, including a four-chambered heart.

Most mammals belong to the placental group. The four largest orders within the placental mammals are Rodentia (mice, rats, and other small, gnawing mammals), Chiroptera (bats), Carnivora (dogs, cats, bears, and other mammals that primarily eat meat), and Cetartiodactyla (including numerous herbivore species, such as deer, sheep, goats, and buffalos, plus whales). The human species is also a placental mammal, a member of the order Primates.

Distinguishing features

However, other features are required when classifying fossils, since soft tissue glands and some other features are not visible in fossils. Paleontologists use a distinguishing feature that is shared by all living mammals (including monotremes), but is not present in any of the early Triassic synapsids: mammals use two bones for hearing that were used for eating by their ancestors. The earliest synapsids had a jaw joint composed of the articular (a small bone at the back of the lower jaw) and the quadrate (a small bone at the back of the upper jaw). Most reptiles and non-mammalian synapsids use this system including lizards, crocodilians, dinosaurs (and their descendants the birds), and therapsids (mammal-like "reptiles"). Mammals have a different jaw joint, however, composed only of the dentary (the lower jaw bone which carries the teeth) and the squamosal (another small skull bone). In mammals the quadrate and articular bones have become the incus and malleus bones in the middle ear. Note: "non-mammalian synapsids" above implies that mammals are a sub-group of synapsids, and that is exactly what cladistics says they are.

Mammals also have a double occipital condyle: they have two knobs at the base of the skull which fit into the topmost neck vertebra, and other vertebrates have a single occipital condyle. Paleontologists use only the jaw joint and middle ear as criteria for identifying fossil mammals, as it would be confusing if they found a fossil that had one feature, but not the other.

Respiratory system

The lungs of mammals have a spongy texture and are honeycombed with epithelium having a much larger surface area in total than the outer surface area of the lung itself. The lungs of humans are typical of this type of lung.

Breathing is largely driven by the muscular diaphragm at the bottom of the thorax. Contraction of the diaphragm pulls the bottom of the cavity in which the lung is enclosed downward. Air enters through the oral and nasal cavities; it flows through the larynx and into the trachea, which branches out into bronchi. Relaxation of the diaphragm has the opposite effect, passively recoiling during normal breathing. During exercise, the diaphragm contracts, forcing the air out more quickly and forcefully. The rib cage itself also is able to expand and contract to some degree, through the action of other respiratory and accessory respiratory muscles. As a result, air is sucked into or expelled out of the lungs, always moving down its pressure gradient. This type of lung is known as a bellows lung as it resembles a blacksmith's bellows.

Circulatory system

The mammalian heart has four chambers: the right atrium, right ventricle, left atrium, and left ventricle. Atria are for receiving blood; ventricles are for pumping blood to the lungs and body. The ventricles are larger than the atria and their walls are thick, because muscular walls are needed to forcefully pump the blood from the heart to the body and lungs. Deoxygenated blood from the body enters the right atrium, which pumps it to the right ventricle. The right ventricle pumps blood to the lungs, where carbon dioxidediffuses out, and oxygen diffuses in. From the lungs, oxygenated blood enters the left atrium, where it is pumped to the left ventricle (the largest and strongest of the 4 chambers), which pumps it out to the rest of the body, including the heart's own blood supply.

Nervous system

All mammalian brains possess a neocortex, a brain region that is unique to mammals.

Integumentary system

The epidermis is typically ten to thirty cells thick; its main function being to provide a waterproof layer. Its outermost cells are constantly lost; its bottommost cells are constantly dividing and pushing upward. The middle layer, the dermis, is fifteen to forty times thicker than the epidermis. The dermis is made up of many components such as bony structures and blood vessels. The hypodermis is made up of adipose tissue. Its job is to store lipids, and to provide cushioning and insulation. The thickness of this layer varies widely from species to species.

Although mammals and other animals have cilia that superficially may resemble it, no other animals except mammals have hair. It is a definitive characteristic of the order. Some mammals have very little, albeit in obscure parts of their bodies, but nonetheless, careful examination reveals the characteristic. None are known to have hair that naturally is blue or green in color although some cetaceans, along with the mandrills appear to have shades of blue skin. Many mammals are indicated as having blue hair or fur, but in all known cases, it has been found to be a shade of gray. The two-toed sloth and the polar bear may seem to have green fur, but this color is caused by algae growths.

Mammals have sweat glands, a defining feature present only in mammals. Some of these glands produce milk (in what are called mammary glands), a liquid used by newborns as their primary source of nutrition. The monotremes branched from other mammals early on, and do not have the nipples seen in most mammals, but they do have mammary glands.

Physiology

Endothermy

Nearly all mammals are endothermic ("warm-blooded"). Most mammals also have hair to help keep them warm. Like birds, mammals can forage or hunt in cold weather and climates where reptiles and large insects cannot.

Endothermy requires plenty of food energy, so pound for pound mammals eat more food than reptiles. Small insectivorous mammals eat prodigious amounts for their size.

Intelligence

In intelligent mammals, such as primates, the cerebrum is larger relative to the rest of the brain. Intelligence itself is not easy to define, but indications of intelligence include the ability to learn, matched with behavioral flexibility. Rats, for example, are considered to be highly intelligent as they can learn and perform new tasks, an ability that may be important when they first colonize a fresh habitat. In some mammals, food gathering appears to be related to intelligence: a deer feeding on plants has a brain relatively smaller than a cat, who must think to outwit its prey.

Social structure

The dependence of the young mammal on its mother for nourishment has made possible a period of training. Such training permits the nongenetic transfer of information between generations. The ability of young mammals to learn from the experience of their elders has allowed a behavioral plasticity unknown in any other group of organisms and has been a primary reason for the evolutionary success of mammals. The possibility of training is one of the factors that has made increased brain complexity a selective advantage. Increased associational potential and memory extend the possibility of learning from experience, and the individual can make adaptive behavioral responses to environmental change. Individual response to short-term change is far more efficient than genetic response.

Some types of mammals are solitary except for brief periods when the female is in estrus. Others, however, form social groups. Such groups may be reproductive or defensive, or they may serve both functions. In those cases that have been studied in detail, a more or less strict hierarchy of dominance prevails. Within the social group, the hierarchy may be maintained through physical combat between individuals, but in many cases stereotyped patterns of behaviour evolve to displace actual combat, thereby conserving energy while maintaining the social structure.

A pronounced difference between sexes (sexual dimorphism) is frequently extreme in social mammals. In large part this is because dominant males tend to be those that are largest or best-armed. Dominant males also tend to have priority in mating or may even have exclusive responsibility for mating within a “harem.” Rapid evolution of secondary sexual characteristics, including size, can take place in a species with such a social structure.

A complex behavior termed “play” frequently occurs between siblings, between members of an age class, or between parent and offspring. Play extends the period of maternal training and is especially important in social species, providing an opportunity to learn behaviour appropriate to the maintenance of dominance.

Locomotion

Mammals evolved from four-legged ancestors. They use their limbs to walk, climb, swim, and fly. Some land mammals have toes that produce claws and hooves for climbing and running. Aquatic mammals such as whales and dolphins have fins which evolved from legs.

Terrestrial

Specialization in habitat preference has been accompanied by locomotor adaptations. Terrestrial mammals have a number of modes of progression. The primitive mammalian stock walked plantigrade—that is, with the digits, bones of the midfoot, and parts of the ankle and wrist in contact with the ground. The limbs of ambulatory mammals are typically mobile, capable of considerable rotation.

Mammals modified for running are termed cursorial. The stance of cursorial species may be digitigrade (the complete digits contacting the ground, as in dogs) or unguligrade (only tips of digits contacting the ground, as in horses). In advanced groups limb movement is forward and backward in a single plane.

Saltatory (leaping) locomotion, sometimes called “ricochetal,” has arisen in several unrelated groups (some marsupials, lagomorphs, and several independent lineages of rodents). This mode of locomotion is typically found in mammals living in open habitats. Jumping mammals typically have elongate, plantigrade hind feet, reduced forelimbs, and long tails. Convergent evolution within a given adaptive mode has contributed to the ecological similarity of regional mammalian faunas.

Mammals of several orders have attained great size (elephants, hippopotamuses, and rhinoceroses) and have converged on specializations for a ponderous mode of locomotion referred to as “graviportal.” These animals have no digit reduction and deploy the digits in a circle around the axis of the limb for maximum support, like the pedestal of a column.

Arboreal

Well-adapted arboreal mammals frequently are plantigrade, five-toed, and equipped with highly mobile limbs. Some species, including many New World monkeys, have a prehensile tail, which is used like a fifth hand. Brachiation, or “arm walking,” in which the animal hangs from branches and moves by a series of long swings, is an adaptation seen in gibbons. The primitive opposable anthropoid thumb is reduced as a specialization for this method of locomotion. Tarsiers are highly arboreal primates that have expanded pads on the digits to improve grasping, whereas many other arboreal mammals have claws or well-developed nails.

Sloths travel slowly along branches rather than swinging energetically.

Aquatic

Four mammalian groups are fully aquatic. Sirenians (dugongs and manatees) eat plants. Cetaceans (whales, dolphins, and porpoises) and pinnipeds (seals and walruses) eat krill or fish. The sea otter eats a variety of invertebrates and fish. Some semiaquatic mammals are very similar to their close land-borne relatives (otters, muskrats, and water shrews, for example). Other mammals have undergone profound adaptation for swimming and life at sea. Walruses and seals give birth to and nurse their young on land, but cetaceans are completely helpless out of water. They depend on water for mechanical support and thermal insulation. Buoyed by their aquatic environment, whales have evolved into the largest mammals and indeed the largest animals ever.

Aerial

Bats are the only truly flying mammals. Only with active flight have the resources of the aerial habitat been successfully exploited. Mammals belonging to other groups (colugos, marsupials, rodents) are adapted for gliding. A gliding habit is frequently accompanied by scansorial (climbing) locomotion. Many nongliders, such as tree squirrels, are also scansorial.

Feeding

To maintain a high constant body temperature is energy expensive- mammals therefore need a nutritious and plentiful diet. While the earliest mammals were probably predators, different species have since adapted to meet their dietary requirements in a variety of ways. Some eat animalprey- this is a carnivorous diet (and includes insectivorous diets). Other mammals, called herbivores, eat plants. An herbivorous diet includes sub-types such as fruit-eating and grass-eating. An omnivore eats boths prey and plants. Carnivorous mammals have a simple digestive tract, because the proteins, lipids, and minerals found in meat require little in the way of specialized digestion. Plants, on the other hand, contain complex carbohydrates, such as cellulose. The digestive tract of an herbivore is therefore host to bacteria that ferment these substances, and make them available for digestion. The bacteria are either housed in the multichambered stomach or in a large cecum. The size of an animal is also a factor in determining diet type. Since small mammals have a high ratio of heat losing surface area to heat generating volume, they tend to have high-energy requirements and a high metabolic rate. Mammals that weigh less than about 18oz (500g) are mostly insectivorous because they cannot tolerate the slow, complex digestive process of a herbivore. Larger animals on the other hand generate more heat and less of this heat is lost. They can therefore tolerate either a slower collection process (those that prey on larger vertebrates) or a slower digestive process (herbivores). Furthermore, mammals that weigh more than 18oz (500g) usually cannot collect enough insects during their waking hours to sustain themselves. The only large insectivorous mammals are those that feed on huge colonies of insects (ants or termites).

Evolutionary history

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The evolution of mammals from synapsids, also known as mammal-like "reptiles" was a gradual process which took approximately 70 million years, from the mid-Permian to the mid-Jurassic, and by the mid-Triassic there were many species that looked like mammals. Note that synapsids are not reptiles at all, but belong to a distinct lineage of tetrapods.

Evolution

The first fully terrestrial vertebrates were amniotes. Like the amphibians they evolved from, they had legs and lungs. Amniotes' eggs, however, had internal membranes which allowed the developing embryo to breathe but kept water in. This allowed amniotes to lay eggs on dry land, while amphibians generally need to lay their eggs in water.

One synapsid group, the pelycosaurs, were the most common land vertebrates of the early Permian and included the largest land animals of the time.

Therapsids descended from pelycosaurs in the middle Permian, about 260M years ago, and took over their position as the dominant land vertebrates. They differ from pelycosaurs in several features of the skull and jaws, including: larger temporal fenestrae; incisors which are equal in size. The therapsids went through a series of stages, beginning with animals which were very like their pelycosaur ancestors and ending with the Triassic cynodonts, some of which could easily be mistaken for mammals:

progress towards an erect limb posture, which would increase the animals' stamina by avoiding Carrier's constraint. But this process was slow and erratic - for example: all herbivorous therapsids retained sprawling limbs (some late forms may have had semi-erect hind limbs); Permian carnivorous therapsids had sprawling forelimbs, and some late Permian ones also had semi-sprawling hindlimbs. In fact modern monotremes still have semi-sprawling limbs.

there is possible evidence of hair in Triassic therapsids, but none for Permian therapsids.

some scientists have argued that some Triassic therapsids show signs of lactation.

The Permian–Triassic extinction event ended the dominance of the therapsids, and in the early Triassic all the medium to large land animal niches were taken over by archosaurs, which were the ancestors of crocodilians, pterosaurs, dinosaurs and birds. After this "Triassic Takeover" the cynodonts and their descendants could only survive as small, mainly nocturnal insectivores. This may actually have accelerated the evolution of mammals - for example the surviving cynodonts and their descendants had to evolve towards warm-bloodedness because their small bodies would otherwise have lost heat quickly, especially as they were active mainly at night.

The first true mammals appeared in the earlyJurassic, over 70 million years after the first therapsids and approximately 30 million years after the first mammaliaformes. Hadrocodium appears to be in the middle of the transition to true mammal status — it had a mammalian jaw joint (formed by the dentary and squamosal bones, but there is some debate about whether its middle ear was fully mammalian.

they use the same orifice to urinate, defecate and reproduce ("monotreme" means "one hole") - as lizards and birds also do.

they lay eggs which are leathery and uncalcified, like those of lizards, turtles and crocodilians.

Unlike other mammals, female monotremes do not have nipples and feed their young by "sweating" milk from patches on their bellies.

The oldest known marsupial is Sinodelphys, found in 125M-year old early Cretaceousshale in China's northeastern Liaoning Province. The fossil is nearly complete and includes tufts of fur and imprints of soft tissues.

Epipubic bones extending forwards from the pelvis, which are not found in any modern placental, but are found in marsupials, monotremes and mammaliformes such as multituberculates. In other words, they appear to be an ancestral feature which subsequently disappeared in the placental lineage.

A narrow pelvic outlet, which indicates that the young were very small at birth and therefore pregnancy was short, as in modern marsupials. This suggests that the placenta was a later development.

It is not certain when placental mammals evolved - the earliest undisputed fossils of placentals come from the early Paleocene, after the extinction of the dinosaurs.

Mammals and near-mammals expanded out of their nocturnal insectivore niche from the mid Jurassic onwards - for example Castorocauda had adaptations for swimming, digging and catching fish.

The traditional view is that: mammals only took over the medium- to large-sized ecological niches in the Cenozoic, after the extinction of the dinosaurs; but then they diversified very quickly; for example the earliest known bat dates from about 50M years ago, only 15M years after the extinction of the dinosaurs.

On the other hand recent molecular phylogenetic studies suggest that most placental orders diverged about 100M to 85M years ago, but that modern families first appeared in the late Eocene and early Miocene But paleontologists object that no placental fossils have been found from before the end of the Cretaceous

During the Cenozoic several groups of mammals appeared which were much larger than their nearest modern equivalents - but none was even close to the size of the largest dinosaurs with similar feeding habits.

Earliest appearances of features

Hadrocodium, whose fossils date from the early Jurassic, provides the first clear evidence of fully mammalian jaw joints.

It has been suggested that the original function of lactation (milk production) was to keep eggs moist. Much of the argument is based on monotremes (egg-laying mammals):

The earliest clear evidence of hair or fur is in fossils of Castorocauda, from 164M years ago in the mid Jurassic. From 1955 onwards some scientists have interpreted the foramina (passages) in the maxillae (upper jaws) and premaxillae (small bones in front of the maxillae) of cynodonts as channels which supplied blood vessels and nerves to vibrissae (whiskers), and suggested that this was evidence of hair or fur. But foramina do not necessarily show that an animal had vibrissae - for example the modern lizard Tupinambis has foramina which are almost identical to those found in the non-mammalian cynodont Thrinaxodon.

The evolution of erect limbs in mammals is incomplete — living and fossil monotremes have sprawling limbs. In fact some scientists think that the parasagittal (non-sprawling) limb posture is a synapomorphy (distinguishing characteristic) of the Boreosphenida, a group which contains the Theria and therefore includes the last common ancestor of modern marsupial and placentals - and therefore that all earlier mammals had sprawling limbs. Sinodelphys (the earliest known marsupial) and Eomaia (the earliest known eutherian) lived about 125M years ago, so erect limbs must have evolved before then.

It is currently very difficult to be confident when endothermy first appeared in the evolution of mammals. Modern monotremes have a lower body temperature and more variable metabolic rate than marsupials and placentals. So the main question is when a monotreme-like metabolism evolved in mammals. The evidence found so far suggests Triassiccynodonts may have had fairly high metabolic rates, but is not conclusive. In particular it is difficult to see how small animals can maintain a high and stable body temperature without fur, and there is no certain evidence of fur before Castorocauda, about 164M years ago.

Classification

George Gaylord Simpson's "Principles of Classification and a Classification of Mammals" (AMNH Bulletin v. 85, 1945) was the original source for the taxonomy listed here. Simpson laid out a systematics of mammal origins and relationships that was universally taught until the end of the 20th century. Since Simpson's classification, the paleontological record has been recalibrated, and the intervening years have seen much debate and progress concerning the theoretical underpinnings of systematization itself, partly through the new concept of cladistics. Though field work gradually made Simpson's classification outdated, it remained the closest thing to an official classification of mammals.

Standardized textbook classification

A somewhat standardized classification system has been adopted by most current mammalogy classroom textbooks. The following taxonomy of extant and recently extinct mammals is from Vaughan et al. (2000).

The McKenna/Bell hierarchical listing of all of the terms used for mammal groups above the species includes extinct mammals as well as modern groups, and introduces some fine distinctions such as legions and sublegions (ranks which fall between classes and orders) that are likely to be glossed over by the nonprofessionals.

The published re-classification forms both a comprehensive and authoritative record of approved names and classifications and a list of invalid names.

Molecular classification of placentals

Molecular studies based on DNA analysis have suggested new relationships among mammal families over the last few years. Most of these findings have been independently validated by Retrotransposonpresence/absence data. The most recent classification systems based on molecular studies have proposed four groups or lineages of placental mammals. Molecular clocks suggest that these clades diverged from early common ancestors in the Cretaceous, but fossils have not been found to corroborate this hypothesis. These molecular findings are consistent with mammal zoogeography:

Following molecular DNA sequence analyses, the first divergence was that of the Afrotheria 110–100 million years ago. The Afrotheria proceeded to evolve and diversify in the isolation of the African-Arabian continent. The Xenarthra, isolated in South America, diverged from the Boreoeutheria approximately 100–95 million years ago. According to an alternative view, the Xenarthra has the Afrotheria as closest allies, forming the Atlantogenata as sistergroup to Boreoeutheria. The Boreoeutheria split into the Laurasiatheria and Euarchontoglires between 95 and 85 mya; both of these groups evolved on the northern continent of Laurasia. After tens of millions of years of relative isolation, Africa-Arabia collided with Eurasia, exchanging Afrotheria and Boreoeutheria. The formation of the Isthmus of Panama linked South America and North America, which facilitated the exchange of mammal species in the Great American Interchange. The traditional view that no placental mammals reached Australasia until about 5 million years ago when bats and murine rodents arrived has been challenged by recent evidence and may need to be reassessed. These molecular results are still controversial because they are not reflected by morphological data, and thus not accepted by many systematists. Further there is some indication from Retrotransposon presence/absence data that the traditional Epitheria hypothesis, suggesting Xenarthra as the first divergence, might be true.